Hydrangea oerstedii Briq., Annuaire Conserv. Jard. Bot. Geneve 20: 407-408. 1919.

Samain, Marie-Stephanie, Granados Mendoza, Carolina & Martinez Salas, Esteban Manuel, 2021, On Hydrangea peruviana, an endangered species from Ecuador, and Hydrangea oerstedii, very common in Costa Rica and Panama, and seven threatened Central and South American Hydrangeas, which have been confounded with these, PhytoKeys 171, pp. 91-153 : 91

4. Hydrangea oerstedii Briq., Annuaire Conserv. Jard. Bot. Geneve 20: 407-408. 1919. Figures 7 , 8

Cornidia radiata Oerst., Vidensk. Meddel. Naturhist. Foren. Kjobenhaven 1856: 42. 1856. Type. Costa Rica. Cartago: in monte Candelaria, 6000-7000 ft, Feb 1847, ♂, flowers, A.S. Oersted 1782 (lectotype, designated by McClintock 1957, pg. 238 [as “holotype”]: C!, isolectotypes: F! [V0066575F], US! [00097007]).

Hydrangea peruviana var. oerstedii (Briq.) Freire Fierro, Fl. Ecuador 73: 34. 2004. Type. Based on Hydrangea oerstedii Briq.

Type.

Based on (replacement name for) Cornidia radiata Oerst.

Description.

Root-climbing liana of up to 60 m high, often bending downwards, functionally dioecious; free-growing branches quadrangular, glabrous (Fig. 7A ); leaves decussate, petiole flattened or sulcate adaxially, terete abaxially, color dark green, pubescent with caducous, appressed, white stellate hairs, 1.5-3 cm long, leaving a rhomboid scar on the branch when leaves fall; lamina flat, ovate to lanceolate-elliptic, 8-25 cm long, 4-14 cm broad, base cuneate, apex acuminate, leaf margin glandular-dentate, venation brochidodromous, veins 6-9 pairs, adaxial leaf side with midvein and primary veins sunken, secondary veins slightly visible, primary veins joining to form a submarginal or intramarginal vein, pubescent with caducous, appressed, white, stellate hairs, older leaves glabrous, abaxially with protruding veins, secondary veins connecting the primary veins, opaque olive green, densely pubescent with caducous, appressed, white, stellate hairs, acarodomatia numerous, consisting of simple cavities, in axils of midvein and primary veins; inflorescence axis angular, pubescent with caducous, appressed, white, stellate hairs, 8-42 cm long, with 3-6 decussate leaf below the inflorescence, persistent, petiole 1-1.5 cm long, lamina 6-13 cm long, 2.5-8 cm broad, apex of the floral axis woody, cone-shaped, elongated bract scars visible, thickening at the top, 6-11 mm broad, 3-7 mm high in functionally female plants, 10 mm broad, 5 mm high in functionally male plants, inflorescence bracts cucullate, cream-colored, coriaceous, abaxial surface tubercled, margin pubescent, adaxially white pubescent, margin membranous, veins slightly marked, bracts increasing in size, lowermost bract 3 cm large, 4 cm broad, higher bracts up to 5 cm large, 5.5 cm broad, consecutively and rapidly deciduous during inflorescence development, inflorescences lateral, decussate, 1-4 pairs of inflorescences per flowering branch (Fig. 7B ), sometimes only one inflorescence developing, flowering branch continues growing vegetatively with the same vigor and periodicity above the inflorescences as below these, inflorescence axes with basal lignified parts of inflorescences of previous years visible in well-collected specimens, allowing to observe growth and flower periodicity, these rests 18-51 cm apart, with 5-11 decussate leaf pairs, but constant within one specimen (Samain et al. 2012-047, Samain et al. 2012-072), medulla central in the branch, disappearing in older branches, leaving a hole only, inflorescence umbellate, buds up to 4-5 cm broad and 3.5-4 cm high before opening, in flowering stage 6-28 cm diameter, 4-17 cm high, with 8 main axes in functionally male plants (Fig. 7C, D ), 5-14 main axes in functionally female plants (Fig. 7F ), partial inflorescences cymes, lax, secondary and tertiary inflorescence axes towards the exterior glabrous, towards its center densely pubescent with whitish stellate hairs; enlarged marginal flowers generally present (Fig. 7B-D ), rarely absent (e.g. Haber 1511), terminally placed in a cyme, sepals 4, pinkish red to wine red, whitish in old inflorescence and infructescences, separate at their base, with marked veins, palmate, pistils 2 or reduced, central part of the flower often amorph, 1-1.5 cm diameter, growing up to 4 cm diameter during flower development, further characters not observed in detail, pedicel 1-4 cm long; flower pedicel of reduced flowers (0.2-)0.5-2(-4) mm long in functionally male flowers, 2-0.5(-2) mm long in functionally female flowers, receptacle broadly triangular in functionally male flowers, semiglobose to semirectangular in functionally female flowers, ovary inferior, calyx lobes 4, triangular, 0.2-0.3 mm long, 0.8 mm broad, petals 4, pinkish red to purple, valvate, cucullate, papyraceous, 1-2 mm long, 0.6-1.0 mm broad; functionally male flowers (Fig. 7E ): hypanthium 1.5-2.0 mm diameter, stamens 8, well-developed, filaments 1.5-3 mm long, anthers 0.8-1.2 mm long, 0.5-0.8 mm broad, pistils 2, reduced, 0.5 mm long, transparent, stigmas not penicellate; functionally female flowers (Fig. 7F ): hypanthium 1.2-2 mm diameter, 1 mm high, stamens 8, reduced, filaments 0.5 mm long, anthers 0.5 mm long, 0.2 mm broad, pistils 2, 1.2-1.5 mm long, rarely enlarging up to 2 mm during fruit maturation, stigmas apically clavate and shortly penicellate; fruit a semiglobose to semirectangular capsule, apically with a conspicuous, thickened border, ribs 4-6, dark brown with dark reddish brown margin, 1.8-3 mm high, 1.8-3 mm broad above, 1.5-2.5 mm diameter, opening between the two pistils to release seeds, seeds not seen.

Distribution.

Hydrangea oerstedii is very abundant in Costa Rica and Panama. It is the most common species of this study (Fig. 8 ).

Habitat.

This species occurs in mountain cloud forest at elevations between 1100 and 2500 m. In contrast to most of the other Hydrangea species, it is common to find this species in disturbed habitats, even when there is no primary forest left and only the very humid microhabitat and protection against direct sunlight remain.

Phenology.

This species has been collected with flowers and fruits throughout the year.

Notes.

Not to be considered as a variety of H. peruviana , from which it can be distinguished by the flat, ovate to lanceolate-elliptic leaves with white pubescence abaxially and a glandular dentate margin. In contrast, H. peruviana is characterized by very slightly spoon-shaped, elliptic to slightly obovate leaves with reddish pubescence adaxially and a serrate to slightly dentate margin. Moreover, H. oerstedii is a relatively common species in Costa Rica and Panama, whereas H. peruviana is a very rare species from Ecuador.

Freire Fierro (2004) also mentioned the presence of an isotype in K, but although this specimen bears the same locality data on its label as the type, it cannot be confirmed that it concerns the same collection, given that no number is mentioned.

According to our molecular study, Hydrangea oerstedii is not related to any of the species included in this study (Granados Mendoza et al. unpublished results).

Preliminary conservation status.

With an EOO of slightly more than 20,760 km2 and an AAO of 428 km2, this species is Near Threatened according to the IUCN categories and criteria ( IUCN 2012). Although its habitat is continuously being destroyed and fragmented, this species is robust, and basically only needs a host tree that remains of the primary forest. As the older plants bend downwards and often have their eye-catching inflorescences just a few meters above ground level, it is even possible that this species is favored when trees are being logged. Nevertheless, should large areas of forest be destroyed, this species may well enter into a threatened category.

Additional specimens examined.

Costa Rica. Alajuela: Alajuela, Distrito 3° Carrizal, 13.1 km N of Heredia via Rte. 114 (10.9 km. N of Barva), 20 ft. from rd., 10°5'26.4"N, 84°9'56.71"W, 1500-2000 m, 15 May 1990, ♀, fruits, Grantham 90.0433 & Parsons (CAS); San Ramón, Distrito 1° San Ramón, Vicinity of San Ramón, La Palma de San Ramón, 14 Dec 1926, ♀, fruits, Brenes 5221 (CR, NY); R.B. Monteverde, Cordillera de Tilarán, Sendero Rancho Alegre, 10°17'40"N, 84°45'W, 1000 m, 4 Apr 1995, ♀, fruits, Martínez 379 (CR, MO); La Palma, Picada San Antonio, 1 Dec 1922, ♀, fruits, Brenes 3807 (CR, F, GH); Reserva Biológica Bosque Nuboso de Monteverde, al lado del Sendero Chomogo, 1.15 km al NE de la caseta de entrada, 10°18'22.23"N, 84°47'11.12"W, 1710 m, 2 Aug 2012, ♀, fruits, Samain & Martínez 2012-045bis (CR, GENT, IEB, MEXU); Peñas Blancas, R.B. Monteverde, Peñas Blancas River Valley, Atlantic slope, 10°20'N, 84°50'W, 1250 m, 12 Oct 1985, ♂, flowers, Haber 3102 (CR); Reserva Biológica Bosque Nuboso de Monteverde, al lado del Sendero Camino, 10°17'59.87"N, 84°46'58.35"W, 21 Jan 2013, 1519 m, ♂, flowers, Samain & Martínez 2013-025 (CR, GENT, IEB, MEXU); cerca de División continental (Ventana), 1550-1580 m, 14 Dec 1976, ♀, flower buds, fruits, Dryer 1080 (CR, F, MO); Monteverde Reserve, Atlantic Slope, Río Peñas Blancas valley, ridge trail above Quebrada Leona, 1050-1300 m, 17 Jan 1985, ♀, flowers, fruits, Haber 1304 (MO); Monteverde Reserve, Atlantic Slope, Río Peñas Blancas valley, 960 m, 19 Nov 1984, ♀, fruits, Haber 1035 (MO); same data as preceding, 1300 m, 20 Mar 1985, ♀, flower buds, Haber 1511 (MO); Reserva Biológica Monteverde, Río Peñas Blancas, Parcela de Rockwell, 10°19'N, 84°43'W, 820 m, 21 Dec 1988, ♂, flowers, Bello 632 (CR, MO); Alajuela, no protegida, Cuenca del Sarapiquí, Hotel Posada Volcán Poás, Viento fresco, 10°10'27.81"N, 84°10'10.072"W, 1800 m, 13 Mar 2003, ♀, old fruits, Kriebel 2911 (CR); San Ramón, Cerros de la Palma, 1500 m, 24 Jan 1984, ♀, fruits, Gómez-Laurito 9802 (CR, F); San Ramón, La Palma, 13 Mar 1929, ♀, fruits, Brenes 19368 (CR); La Palma, Picada de San Antonio, Dec 1922, ♀, old fruits, Brenes 19366 (CR); La Palma de San Ramón, 22 Dec 1922, ♀, old fruits, Brenes 19367 (CR); Bosque Húmedo en San Pedro de San Ramón, 11 Jan 1924, ♀, old fruits, Brenes 19370 (CR); La Palma, Picada de San Antonio, Dec 1922, ♀, old fruits, Brenes 19371 (CR); San Ramón, Bosque húmedo de las colonias de San Pedro, 1200 m, 11 Jan 1924, ♀, fruits, Brenes 3984 (F); Cartago: Cartago, Distrito 5° San Francisco, 0.5 km N. of Tapantí, 9°47'56.64"N, 83°55'37.57"W, 1300 m, 7 Nov 1971, ♀, young fruits, Lent 2210 (F, GB, US); El Guarco, Distrito 2° San Isidro, dry wooded slopes about 13 km south of the Taras Intersection along the Interamerican Highway or about 18 km north of El Empalme, 9°47'52.05"N, 83°56'49.10"W, 4 Jul 1976, ♂, old flowers, Wilbur 19852 (DUKE); Empalme, Cartago, 9°43'5.91"N, 83°56'50.32"W, 2300 m, 1 Dec 1983, ♂, flower buds, flowers, Gómez 22176 (CR, F, MO, UC); Cartago, slopes in the vicinity of Palo Verde and the Quebrada Cangreja on the road to Estrella, 9°47'0.07"N, 83°57'11.48"W, 1600 m, 19 Dec 1974, ♀, fruits, Wilbur 18437 & Luteyn (CAS, DUKE, F); Turrialba, Distrito 8° Tayutic, Turrialba, Jicotea, por la fila al lado oeste del Río Jicotea, Finca del señor Israel Martínez, 9°47'5"N, 83°33'15"W, 1100-1200 m, 7 Dec 1994, ♀, fruits, Sánchez et al. 424 (CR, F, K); Cartago, hills above Platanillo, 9°49'31.24"N, 83°24'16.91"W, 915 m, 7 Nov 1967, ♂, old flowers, Stone 2324 (DUKE); Paraíso, Distrito 3° Orosi, about 15 km south of Tapantí along the new road, on the east slope above the Río Grande de Orosi near the concrete bridge, 9°46'44.28"N, 83°47'46.78"W, 1500 m, 30 Dec 1970, ♀, fruits, Burger 7525 & Liesner (F, GB); Cartago, Distrito 5° San Francisco, about 3-5 km S of Muñeco along the Río Sombrero, 9°45'38.34"N, 83°54'21.62"W, 1400-1500 m, 25 Feb 1978, ♀, fruits, Wilbur 24976 (DUKE); Cartago, ca. 2.5 km S of Muñeco, 9°46'15.95"N, 83°54'17.34"W, 1500 m, 25 Feb 1978, ♀, fruits, Almeda & Nakai 3937 (CAS, CR); El Guarco, Distrito 2° San Isidro, Alto de La Estrella, 26-27 Mar 1924, ♀, fruits, Standley 39080 (US); Cartago, al lado del camino Copatchi-San Rafaël Arriba, 9°49'47.52"N, 84°2'22.83"W, 1838 m, 27 Jul 2012, sterile, Samain & Martínez 2012-034 (CR, GENT, IEB, MEXU); al lado del camino Copatchi-Quebradilla, 9°49'54.96"N, 84°1'38.37"W, 1931 m, 27 Jul 2012, sterile, Samain & Martínez 2012-035 (CR, GENT, IEB, MEXU); El Guarco, Cuenca del Reventazón, San Isidro, Ruta al Cerro de la Muerte, entrada a Palo Verde, 1 km después de la entrada, 9°47'56"N, 83°56'51"W, 1800 m, 26 Nov 1998, inflorescence buds, Rodríguez et al. 4208 (CR, MO); camino San Isidro de El General-Cartago, 9°41'23.46"N, 83°54'12.64"W, 2532 m, 11 Aug 2012, ♀, fruits, Samain & Martínez 2012-072 (CR, GENT, IEB, MEXU); camino Cartago-San Isidro de El General, 9°46'16.46"N, 83°59'1.03"W, 1770 m, 23 Jan 2013, ♂, flowers, Samain & Martínez 2013-030 (CR, GENT, IEB, MEXU); same data as preceding, 9°46'16.46"N, 83°59'1.03"W, 1769 m, 23 Jan 2013, ♀, fruits, Samain & Martínez 2013-031 (CR, GENT, IEB, MEXU); same data as preceding, 9°41'23.72"N, 83°54'12.58"W, 2489 m, 23 Jan 2013, ♀, fruits, Samain & Martínez 2013-032 (CR, GENT, IEB, MEXU); Paraíso, P.N. Tapantí, Valle del Reventazón, Río Grande de Orosi, fila innominada al sur del cruce de la toma del Río Humo, 9°43'5"N, 83°46'50"W, 1500-1600 m, 21 Jun 1996, floral buds, Morales 5394 et al. (CR, MO); Cartago, Paraíso, Cordillera de Talamanca, Río Grande de Orosí, Tapantí, bosque alterado por crecientes del Río Villegas, cerca de la toma de agua, 9°41'50"N, 83°46'50"W, 1500 m, 22 Jul 1994, ♂, old flowers, Morales 3059 (CR, F); Cartago, El Guarco, No protegida, Cuenca del Pirrís-Damas, Cerros de la Muerte, 9°47'47.6085"N, 83°59'54.3782"W, 2100 m, 18 Feb 2002, ♂, young flowers, González 1528 (CR); Cartago, Paraíso, P.N. Tapantí, Valle del Reventazón, Estación Tapantí, Sendero La Heliconia, 9°45'20"N, 83°47'W, 1300 m,10 Jun 1995, ♂, old flowers, Mora 661 (CR); Cartago, Paraíso, P.N. Tapantí-Macizo de la Muerte, Cuenca del Reventazón, P.N. Tapantí, Sendero Árboles Caídos, 9°45'4"N, 83°47"W, 1300-1413 m, 18 Jun 2005, ♂, old flowers, Morales 13225 (CR); Cartago, Paraíso, No protegida, Cuenca del Reventazón, entrada a Monte Sky, 9°44'53.9065"N, 83°50'4.8525"W, 1537 m, 5 Dec 2000, ♀, fruits, González 1225 (CR); Cartago, Paraíso, P.N. Tapantí, Valle del Reventazón, Sector Dos Amigos, Sendero Rancho Negro, 9°45'20"N, 83°47'W, 1300 m, 20 Jul 1994, floral buds, Morales 289 (CR); Cartago, La Unión, San Rafael, Cerros de La Carpintera, Campo Iztarú de los Scouts, Alrededores del área administrativa, 9°53'24.8"N, 83°58'6.8"W, 1730 m, 13 Oct 2010, ♂, old flowers, Cascante 2250 (CR); Prov. de Cartago, La Unión, Tres Ríos, 1350 m, 12 Apr 1953, ♀, fruits, Córdoba 327 (CR): Cartago, San Nicolás, Cerros de la Carpintera, Campo Escuela Iztarú, propiedad de la Asociación Scouts de Costa Rica, sendero hacia la cima de los cerros, 9°52'55.8"N, 83°58'40.3"W, 1854 m, 11 Jun 2007, ♀, fruits, Estrada 3977 (CR); Cartago, Paraíso, Orosí, Camino del pueblo de Río Macho al Embalse, área de cafetales y pequeños bosques alterados, 9°46'15"N, 83°51'15"W, 1500 m, 26 Jan 2010, ♀, fruits, Cascante 2179 (CR); Tapantí, 1300 m, 15 Jul 1937, floral buds, Valerio 1658 (F); Cartago, very steep slopes with open forest and many epiphytes about 10 km south of Tapantí along the new road on the east slope above th Río Grande de Orosí, 9°42'N, 83°47'W, 1400-1600 m, 10-24 Jun 1968, ♀, fruits, Burger & Stolze 6108 (F); Turrialba, Las Cortinas 1 km al norte del Monumento Guayabo, Manglares del Río Guayabo, 9°57'50"N, 83°41'30"W, 1100 m, 13 Oct 1992, ♀, fruits, Rivera 2007 (CR, F, MO); Refugio Nacional de Vida Silvestre Tapantí, about 7-15 km beyond the main entrance to the reserve; roadsides and trails through wet montane rainforest, 1500-1650 m, 3 Mar 1991, ♀, fruits, Almeda et al. 6912 (CR, MO); Guanacaste: Tilarán, Área no protegida, Cordillera de Tilarán, Reserva Biológica del Colegio de Monteverde, 10°21'20"N, 84°49'50"W, 1600-1700 m, 10 Jan 1992, ♀, fruits, Bello 4328 (CR, MO); Z.P. Tenorio, Cordillera V Tilarán, Tierras Morenas, Río San Lorenzo, 1050 m, 10°36'40"N, 84°59'45"W, 10 May 1994, ♂, old flowers, Rodríguez 13 (CR, MO); Santa Elena, cerca de la entrada del Parque Selvatura, 10°20'31.23"N, 84°47'56.94"W, 1618 m, 1 Aug 2012, ♂, Samain & Martínez 2012-044 (CR, GENT, IEB, MEXU); Santa Elena, en el límite de la Reserva Bosque Nuboso Santa Elena, 10°20'35.12"N, 84°47'50.53"W, 1659 m, 31 Jul 2012, ♀, fruits, Samain & Martínez 2012-045 (CR, GENT, IEB, MEXU); Reserva Bosque Nuboso Santa Elena, al lado del camino a la Laguna de Arenal, 827 m al NE de la caseta de entrada, 10°20'58.70"N, 84°47'33.56"W, 1582 m, 31 Jul 2012, ♀, fruits, Samain & Martínez 2012-046 (CR, GENT, IEB, MEXU); Reserva Bosque Nuboso Santa Elena, al lado del camino a la Laguna de Arenal, 1.27 km al NE de la caseta de entrada, 10°21'14.67"N, 84°47'34.77"W, 1555 m, 31 Jul 2012, ♀, old inflorescence axes, fruits, Samain & Martínez 2012-047 (CR, GENT, IEB, MEXU); Reserva Bosque Nuboso Santa Elena, al lado del camino a la Laguna de Arenal, 415 m al NE de la caseta de entrada, 10°20'48.66"N, 84°47'40.36"W, 1643 m, 31 Jul 2012, ♀, fruits, Samain & Martínez 2012-048 (CR, GENT, IEB, MEXU); al lado del camino de Santa Elena al Mirador Lodge, 1.01 km desde la desviación hacia la Reserva Bosque Nuboso Santa Elena, 10°21'9.35"N, 84°48'6.97"W, 1545 m, 20 Jan 2013, inflorescence buds, floral buds, Samain & Martínez 2013-019 (CR, GENT, IEB, MEXU); al lado del camino de Santa Elena al Mirador Lodge, 1.74 km desde la desviación hacia la Reserva Bosque Nuboso Santa Elena, 10°21'32.08"N, 84°48'9.79"W, 1495 m, 20 Jan 2013, ♀, inflorescence buds, flowers, Samain & Martínez 2013-021 (CR, GENT, IEB, MEXU); al lado del camino de Santa Elena al Mirador Lodge, 2,069 m desde la desviación hacia la Reserva Bosque Nuboso Santa Elena, 10°21'43.64"N, 84°48'16.32"W, 1509 m, 20 Jan 2013, ♀, fruits, Samain & Martínez 2013-022 (CR, GENT, IEB, MEXU); Heredia, Heredia, Distrito 6° San José de la Montaña, Cantón de Barva. Cerca de Porrosatí, 10°6'N, 84°7'W, 2000 m, 6 Jan 1991, ♂, flowers, Hammel 18228 (CR, K, MO); Heredia: Distrito Varablanca, Vara Blanca de Sarapiquí, north slope of Central Cordillera, between Poás and Barba volcanoes, 10°11'2.85"N, 84°9'12"W, 1680 m, 1 Jan 1938, ♀, flowers, fruits, Skutch 3393 (K, MO, US); Region between San Rafaél and the Río San Rafaél about 3 km east of Vara Blanca, 1800 m, 10°10'17.96"N, 84°8'21.84"W, 23 Feb 1976, ♀, flower buds, flowers, fruits, Utley & Utley 4166 (DUKE, MO); Barva, No protegida, Cuenca del Tárcoles, entre Porrosatí y Sacramento, faldas del Volcán Barva, 10°5'22"N, 84°6'22"W, 1950 m, 24 Apr 2002, ♀, young fruits, Hammel 22523 (CR); Barva, P.N. Braulio Carrillo, S slope of west forest, 1950 m, 16 Jan 1965, ♀, flower buds, flowers, Hatheway 1275 (CR); Barva, Montaña, La Isla, Sacramento, 2500 m, ♂, old flowers, 13 Jan 1987, Gómez-Laurito 11304 (CR, F); Barva, Cordillera Central, entre Porrosatí y Sacramento, faldas del Volcán Barva, 10°5'22"N, 84°6'22"W, 2000 m, 24 Jan 1996, ♂, flowers, Hammel 20131 (CR, MO); cord. V. Central, faldas de V. Barva, Porrosatí, camino al volcán Barva, 10°5'45"N, 84°7'10"W, 2000 m, 12 Nov 1993, ♂, flowers, Jiménez 1406 (CR, MO); faldas del Volcán Barva, entre Porrosatí y Sacramento, 2047 m, 10°5'38.20"N, 84°6'12.75"W, 6 Aug 2012, ♀, fruits, Samain & Martínez 2012-041 (CR, GENT, IEB, MEXU); same data as preceding, 10°5'38.28"N, 84°6'15.92"W, 2035 m, 28 Jul 2012, inflorescence buds, Samain & Martínez 2012-042 (CR, GENT, IEB, MEXU); Vara Blanca, terreno del restaurante Vara Blanca, 10°9'32.40"N, 84°9'20.28"W, 1842 m, 15 Jan 2013, ♂, inflorescence buds, flowers, Samain & Martínez 2013-001 (CR, GENT, IEB, MEXU); faldas del Volcán Barva, entre Porrosatí y la entrada al parque, 10°7'17.65"N, 84°7'23.48"W, 2447 m, 18 Jan 2013, ♀, flowers, Samain & Martínez 2013-012 (CR, GENT, IEB, MEXU); Heredia, 13.1 km N of Heredia via Rte. 114 (10.9 km N of Barva), 20 ft from road, 1500-2000 m, 15 May 1990, Grantham & Parsons 0013-90 (CAS); Limón: Talamanca, Cordillera de Talamanca, Quebrada Kuisa, al paso del Sendero entre Ujarrás y San José Cabécar, 9°20'N, 83°13'55"W, 2100 m, 13 Mar 1993, ♀, fruits, Fernández 653 (CR, MO); P.N. La Amistad, cuenca del Sixaola, sobre transecto 5, 9°21'18.1080"N, 83°13'29.9820"W, 1848 m, ♀, fruits, 25 Feb 2007, Solano 4048 (PMA); Parque Internacional La Amistad, sendero Transtalamancana, entre Ujarras y San José Cabécar, alrededores del Río Lori, 9°21'18.6080"N, 83°13'19.2000"W, 1850-1900 m, 25 Feb 2007, ♀, fruits, Rodríguez et al. 10829 (CR, PMA); Talamanca, P.N. La Amistad, Cuenca del Sixaola, transecto 5, 9°21'18.108"N, 83°13'29.892"W, 1848 m, 25 Feb 2007, ♀, fruits, Solano 4046 (CR); Guápiles, Los Angeles, San Miguel, siguiendo el camino que lleva a San Gerardo de Oreamuno, volcán Irazú, 10°6'10"N, 83°50'45"W, 1300 m, 23 Feb 1990, ♂, flowers, Herrera 3789 & Schik (CR, MO); Cordillera de Talamanca, headwaters of the unnamed western branch of the Río Teribe, between the Río Sini and the continental divide at Cerro Bekom, 9°10'45"N, 83°3'30"W, 2500-2600 m, 21&27 Mar 1984, inflorescence bud, floral buds, Davidse et al. 26158 (CR, MO); Puntarenas: Distrito Monteverde, Z. P. Arenal-Monteverde, Cuencas del Lagarto y Guacimal, Reserva Biológica Monteverde, a 1 km del inicio del Sendero principal, 10°19'17"N, 84°48'35"W, 1540 m, 1 Jun 1999, ♀, flower buds, flowers, Hurtado 156 (CR, NY); on and near Continental Divide about 2 to 5 km. east and southeast of Monteverde, 10°18'9.94"N, 84°47'44.07"W, 1580-1700 m, 17-20 Mar 1973, ♀, fruits, Burger 8613 & Gentry, Jr. (DUKE, F, MO); Buenos Aires, P.N. La Amistad, Cuenca Térraba-Sierpe, sendero de Bekon a Tres Colinas, 9°9'42.965"N, 83°4'1.44"W, 2400-2500 m, 18 Apr 2002, ♂, flowers, Alfaro 3978 et al. (CR, MO); Z.P. Las Tablas, Cuenca Térraba-Sierpe, Zona Protectora Las Tablas, Surá, 7 km NO de Progreso, 8°54'17"N, 82°45'3"W, 2050 m, 18 Jan 1987, ♂, old flowers, Navarro 579 (CR, MO); Puntarenas, Reserva Biológica Monteverde, Ojo de Agua, Finca de Leonel Hernández, lado pacífico de la reserva, 10°15'N, 84°46'W, 1600 m,18 Nov 1987, ♀, flower buds, flowers, Haber 7811 & Bello (CR, MO); Monteverde Community, Pacific slope, 1450 m, 10°20'N, 84°50'W, 13 Aug 1986, ♂, old flowers, Haber ex Bello 5295 (MO); Reserva Biológica Monteverde, Pacific slope and continental divide, 10°18'N, 84°48'W, 1600 m, 17 Jun 1992, ♀, fruits, Haber & Zuchowski 11210 (CR, MO); R.B. Monteverde, Cordilla de Tilarán, Monteverde, camino hacia Alto Cebadilla, 10°19'30"N, 84°51'W, 1300 m, 9 Feb 1994, ♀, fruits, Ramírez 242 & Poveda (CR, MO); Monteverde Cloud Forest Reserve, Cordillera de Tilarán, pacific slope of continental divide, Pantanoso trail, 1500-1620 m, 29 Dic 1984, ♀, fruits, Pounds 379 (MO); Forest along trail between Las Alturas and Cotonsito, along the Río Cotón, 8°56'30"N, 82°48'W, 1400 m, 31 Aug 1983, ♂, flowers, Davidse 24651 (CR, F, MO, MEXU); Foothills of the Cordillera de Talamanca, lower montane forest along the Río Bella Vista, NW of Las Alturas, 8°57'N, 82°51'W, 1450-1600 m, 30 Aug 1983, ♂, inflorescence buds, flowers, Davidse 24300 (CR, MO); Monteverde Cloud forest Reserve, continental divide near La Ventana and elfin forest, 10°20'N, 84°50'W, 1600 m, 14 Dec 1985, ♂, inflorescence buds, flowers, Haber ex Bello 3959 (CR, MO); same data as preceding, ♂, flowers, Haber ex Bello 3969 (CR, MO); forest and forest edges on and around Wilson’s finca, 6 km south of San Vito de Java, about 4000 ft, 16 Aug 1967, floral buds, Raven 21806 (F, MO); Puntarenas, Coto Brus, Z.P. Las Tablas, Cuenca Terraba-Sierpe, Sendero Sura, Zonas Protectora Las Tablas, 8°56'57"N, 82°44'37"W, 1920 m, 20 Dec 1997, ♂, old flowers, Navarro 827 (CR, MO); Puntarenas, Puntarenas, Reserva Biológica Monteverde, Ojo de Agua, Finca de Leonel Hernández, lado pacífico de la reserva, 10°15'N, 84°46'W, 1600 m, 18 Nov 1987, floral buds, Haber 7811 & Bello (CR, MO); Puntarenas, Bob Wilson´s Forest, San Vito de Java, near the Panamanian border, 22 Aug 1965, ♂, old flowers, Walker 164 (F); Puntarenas, near Bob Wilson’s finca at San Vito Vine, 20 Aug 1965, ♂, old flowers, Croat 1067 (MO); Foothills of the Cordillera de Talamanca, lower montane forest along the Río Bella Vista, NW of Las Alturas, 8°57'N, 82°51'W, 1450-1600 m, 30 Aug 1983, floral buds, Davidse 24281 (MO); Monte Verde, 1500 m, 27 Dec 1984, ♂, inflorescence buds, flower buds, flowers, de Nevers & Charnley 4427 (MO, PMA); San José: Desamparados, Distrito 2° San Miguel, Altos de Tablazo, about 5 km SE of Higuito on Calle Tablazo or about 12 km SE of Desamperados, 9°49'54.82"N, 84°2'28.28"W, 1800-1900 m, 30 Jan 1976, ♂, inflorescence bud, flowers, Utley & Utley 3838 (DUKE, F); Distrito 8° San Cristobal, Cordillera de Talamanca. 4 km. north of El Empalme, 9°44'45.61"N, 83°58'36.99"W, 2070 m, 3 Mar 1986 ♂, flowers, Almeda & Anderson 5298 (CAS); Distrito 9° Rosario, about 12 km. S of Higuito on the slopes of Altos Tablazo, 9°49'53.39"N, 84°2'26.69"W, 1800 m, 20 Feb 1978, ♂, flower buds, flowers, Almeda 3757 & Nakai (CR, CAS); Dota, Distrito 3° Copey, near Finca La Cima, above Los Lotes, North of El Copey, 9°40'15.4"N, 83°54'48.85"W, 2100-2400 m, 21-22 Dec 1925, ♀, flower buds, flowers, Standley 42569 (US); Pérez Zeledón, Distrito 1° San Isidro de El General, roads above San Isidro de Coronado, 10°0'6.90"N, 84°0'21.38"W, 1400-1600 m, 1 Dec 1937-1 Jan 1938, ♀, flowers, Allen 534 (F, GH); Vasquez de Coronado, Distrito 3° Jesús, La Hondura, 10°3'44.72"N, 83°58'55.21"W, 1300-1700 m, 16 Mar 1924, ♀, fruits, Standley 37781 (US); Lisière des forets a L´Alto de la Palma, 10°2'55.35"N, 83°59'22.02"W, 1542 m, 30 Aug 1898, ♀, fruits, Tonduz 12507 (US); La Palma, 10°2'55.71"N, 83°59'18.93"W, 1600 m, 3 Feb 1924, ♀, fruits, Standley 33163 (US); Río Claro valley ( Río La Hondura drainage) below La Palma northeast of San Jerónimo, 10°3'N, 83°58'W, 19 Nov 1969, 1000-1200 m, inflorescence buds, Burger 6290 & Liesner (CR, F, GB, NY); Woods above Río Hondura, 10°3'30"N, 83°58'28.86"W, 1150 m, 5 Dec 1971, ♀, fruits, Lent 2272 (COL, CR, F, MEXU); heavily wooded slope leading from Alto La Palma to Bajo la Hondura about 12 km NNE of San Vicente de Moravia in Route 220 in the saddle between Irazú and Barba, 10°3'19.33"N, 83°59'10.46"W, 1200 m, 19 May 1971, ♀, fruits, Wilbur 14687 (DUKE); Desamparados, Distrito 2° San Miguel, Altos de Tablazo, about 5 km SE of Higuito on Calle Tablazo or about 12 km SE of Desamparados, 9°49'54.82"N, 84°2'28.28"W, 1800-1900 m, 30 Jan 1976, ♀, fruits, Utley & Utley 3844 (DUKE, F, MO); Perez Zeledon, Distrito, 4° Rivas, P. I. La Amistad, Cordillera de Talamanca, Entre las Nacientes de las quebradas Barranca y Río Blanco, Finca San Carlos, 9°31'47"N, 83°35'30"W, 05 Apr 1995, 2350 m, ♀, fruits, Aguilar 3816 & Garrote (CR, F, K, NY); Dota, San Gerardo de Dota, 2291 m, 9°34'18.56"N, 83°48'4.46"W, 24 Jan 2013, ♂, old flowers, Samain & Martínez 2013-035 (CR, GENT, IEB, MEXU); Escazú, San Antonio, Z.P. Cerros de Escazú, falda SE del Alto Hierbabuena, 9°50'30"N, 84°7'35"W, 2100 m, 15 Oct 1992, ♀, flowers, fruits, Morales 767 (CR, F, MO); Perez Zeledon, al lado del camino a San Gerardo de Dota, 4.22 km al SO del crucero con la Carretera Interamericana San Isidro de El General-Cartago, 9°33'27.20"N, 83°48'14.31"W, 2299 m, 11 Aug 2012, sterile, Samain & Martínez 2012-068bis (CR, GENT, IEB, MEXU); Turrubares, San Luis, Z.P. Cerros Turrubares, faldas del Cerro Bares, 9°47'30"N, 84°28'30"W, 1600 m, 6 Nov 1990, ♀, fruits, Jiménez 919 et al. (CR, F, MO); Bajo de Hondura, P.N. Braulio Carrillo, 10°4'N, 83°58'W, 1100-1200 m, 23 Jan 1983, ♂, old flowers, Davidse 23195 (CR, MO); along Ca-2 on western ascent of Cerro de la Muerte, north of turnoff for road 222, 2000 m, 27 Feb 1976, ♂, inflorescence buds, flowers, Croat 32829 (MO); San José, Sendero del camino viejo, hacia el Río La Hondura, P.N. Braulio Carrillo, 1250 m, 6-9 Dec 1983, ♂, old flowers, Zamora & Carlson 418 (MO); San José, Aserrí, Cerros de Escazú, Tarbaca, bosque secundario en las cabeceras de la quebrada Saurez, 9°50'15"N, 84°7'10"W, 1900-2000 m, 6 Nov 1993, ♀, young fruits, Morales 1979 (CR); San José, Vasquez de Coronado, R.F. Cordillera Volcánica Central, Cuenca del Sarapiquí, Cascajal de Coronado, calle Monserrat, 10°1'4"N, 83°56'31"W, 1700-1800 m, 6 Jul 2000, ♂, old flowers, Vargas 459 (CR); San José, Moravia, No protegida, cuenca del Sarapiquí, paso de la Palma, de las Lecherías, 10°2'50"N, 83°59'20"W, 1500 m, 8 Nov 2003, floral buds, Morales 10053 (CR); San José, Pérez Zeledón, Cordillera de Talamanca, Las Nubes de Santa Elena, tajo a orilla de la carretera, 9°23'30"N, 83°35'50"W, 1210 m, 7 Aug 1995, ♀, fruits, Mora 245 (CR); San José, Parque Nal. Braulio Carrillo, camino entre El Bajo de la Hondura y La Montura, ca. 1200 m, 1 May 1982, ♂, old flowers, Gómez-Laurito 8408 (CR, F); San José, Terrazú, San Lorenzo, R.F. Los Santos, fila San Isidro, 9°34'20"N, 84°4'10"W, 1350 m, 5 Nov 1997, ♂, flowers, Valverde 384 (CR); San José, Dota, Santa María, Cerca de Palmitera de Dota, 1700 m, 20 Dec 1963, ♂, flowers, Jiménez 1466 (CR, F); San José, Dota, Copey, R.F. Los Santos, San Gerardo de Dota, orillas del río Savegre, 2200 m, 26 Nov 1985, ♂, old flowers, Soto s.n. (CR); San José, Pérez Zeledón, Páramo, R.F. Los Santos, Lira, Estribaciones Noroeste de Cerro Lira, 9°31'30"N, 83°51'40"W, 1800 m, 8 Dec 1994, ♂, old flowers, Martén 673 & Herrera (CR, F); San José, P.N. Braulio Carrillo, Sendero del Camino Viejo, hacia El Río La Hondura, 1250 m, 9 Dec 1983, ♂, old flowers, Zamora 417 (CR); without locality, ♂, old flowers, Warcewicz 27368 (MO, photo from Delessert Herbarium).

Panama. Bocas del Toro: Cordillera de Talamanca, headwaters of the Río Culubre, 6 airline km NW of the peak of Cerro Echandi on the Costa Rican-Panamian international border, 09°05'00"N, 82°50'30"W, 2450-2600 m, 2-3 Mar 1984, ♀, fruits, Davidse et al. 25254 (CR, MEXU, MO, PMA); E branch, headwater of Colubre River, 2300-2500 m, 2 Mar 1984, inflorescence bud, Gómez et al. 22303 (MO); La Pata del Cedro, ridgetop, 09°04.270'N, 82°44.174'W, 1750 m, 11 Mar 2004, ♀, fruits, Monro & Alfaro 4315 (CR, MO, PMA); Chiriquí: 1.4 mi S of Cerro Punta, 1850 m, 28 Dec 1963, ♀, fruits, Graham 297 (MICH); along road and in pasturelands and stream banks above Las Nubes, NW of Cerro Punta, 1900-2100 m, 26 May 1973, ♀, fruits, Luteyn 3797 (DUKE, MICH); Parque Internacional La Amistad, 100 m al SSW de la oficina, 8°53'36.3"N, 82°36'58.1"W, 2207 m, 10 May 2019, ♂, inflorescence buds, flowers, Samain et al. 2019 -003 (IEB, MEXU, PMA); Parque Internacional La Amistad, sendero El Retoño, 190 m al WNW de la oficina, 8°53'39"N, 82°37'0.53"W, 2227 m, 10 May 2019, ♀, flowers, Samain et al. 2019 -006 (IEB, MEXU, PMA); along trail between N fork of Río Palo Alto and Cerro Pate Macho, ca 6 km NE of Boquete, 8°48'N, 82°23.5"W, 1600-1700 m, 6 Feb 1986, ♂, old flowers, Grayum et al. 6370 (MO); Alto Pineda, end of road, right turn just before cooperativa entrance to Cerro Punta, 8400 ft, 11 Apr 1979, ♂, inflorescence axes, old flowers, Hammel et al. 6976 (MO); District Boquete, Alto Quiel, Finca Lerida, 05 Nov 1994, Quiroz 166 & Garrido (F, NY, PMA) (old inflorescence axes, flower buds); District Boquete, Bajo Chorro, 6000 ft, 7 Jan 1938, ♀, fruits, Davidson 74 (F, GH, MO, UC, US); edge of forested slope above Cerro Punta toward Bajo Grande in Quebrado Bajo Grande, about 6500 ft, 14 Jan 1970, ♀, flowers, fruits, Wilbur 10923 et al. (DUKE, F, MICH, MO); Jurutungo, de la Fca. Los Quetzales , luego hacia la der. para luego bajar por el Río, 8°54'N, 82°43'W, ca. 1900 m, 24 Sep 1996, ♀, fruits, Aranda 3251 et al. (DUKE, MO, NY, PMA); Las Nubes, 5 km NW of Cerro Punta, 6000-6500 ft, 19 Jul 1975, ♂, flowers, Mori & Bolten 7248 (MO); Monte Azul, 1.4 mi N of Entre Ríos on E slopes of Cerro Punta, 3 mi by road from town to Cerro Punta, 2250 m, 22 Nov 1979, ♀, flower buds, Antonio 2693 (MEXU, MO, PMA); pastured area and bordering woods NE of Cerro Punta about 2.5 km, about 7200 ft, 29 May 1972, ♀, fruits, Wilbur et al. 17149 (DUKE, MICH); road to Cerro Punta National Park from Alto Quiel and Boquete, 8°51'N, 82°29'W, c. 1850 m, inflorescence buds, 16 Jan 1986, McPherson 8031 (MEXU, MO, PMA); trail up Cerro Pate Macho, 08°50'N, 82°25'W, 1500-1900 m, 7 Jan 1983, ♀, fruits, Stein et al. 1215 (MEXU, MO); Valley of the upper Río Chiriquí Viejo, 18 Jan 1938, ♀, fruits, White & White 90a (MO); Vicinity of "New Switzerland", central valley of Río Chiriquí Viejo, 1800-2000 m, 6-14 Jan 1939, ♂, flowers, Allen 1400 (UC, US); 7 km northwest of Cerro Punta, Las Nubes region, 7200 ft, 11 Feb. 1978, floral buds, Hammel 1428 (MO, PMA); vicinity of Fortuna, 8°45'N, 82°15'W, ca. 1250 m, 28 Apr 1986, ♀, fruits, McPherson 9115 (MEXU, MO); Chiriquí, Boquete, Bajo Mono, 1600 m, Jan 1979, ♀, fruits, Beliz 463 (PMA); Boquete, Sendero a Culebra, 1800 m, 18 Jan 2004, sex not visible because of insect damage, Cáceres et al. 1836 (UCH); vicinity of Bajo Mona and Quebrada Chiquero, 1500 m, 18 Jul 1940, ♀, fruits, Woodson & Schery 511 (GH, MO); distrito San Felix, Comarca Gnöbe Buglé, km 0 al 10 entre Hato Chami y Hato Ratón, ± 27-37 km al N de la 8°29'41"N, 81°46'16"W, 1386-1576 m, 20 May 2001, ♀, fruits, Galdames et al. 4691 (F, PMA); Reserva Forestal Fortuna, Sendero La Casa Rosada, bosque cercano al Río Hornito, 8°42'00"N, 82°13'26"W, 1147 m, 31 Jan 2013, ♂, old flowers, Ortiz 1172 et al. (PMA). Without locality, either in Costa Rica or Panama ("Costa Rica et Veragua "), without date, ♂, flowers, Warszewicz 1 (G).

Figures

Figure 7. Hydrangea oerstedii A branch with leaves seen abaxially B flowering branch of functionally male plant with three inflorescences with enlarged marginal flowers and many flowers with large stamens C inflorescence of functionally male plant with enlarged marginal flowers, most flowers in bud with petals visible, and a few open flowers with long stamens D close-up of functionally male inflorescence E close-up of functionally male flowers with reduced pistils F close-up of functionally female flowers with reduced stamens and large pistils. A field image of collection Samain & Martinez 2012 - 044 B field image of collection Samain & Martinez 2013 - 025 C field image of collection Samain & Martinez 2012 - 044 D field image of collection Samain et al. 2019 - 003 E field image of collection Samain & Martinez 2013 - 025 F field image of collection Samain & Martinez 2013 - 021.

Figure 8. Distribution of Hydrangea oerstedii and Hydrangea panamensis.